There is now compelling evidence that microbially mediated reactions impart a significant effect upon the dynamics, composition, and abundance of nutrients in glacial melt water. Consequently, we must now consider ice masses as ecosystem habitats in their own right and address their diversity, functional potential, and activity as part of alpine and polar environments. Although such research is already underway, its fragmentary nature provides little basis for developing modern concepts of glacier ecology. This paper therefore provides a much-needed framework for development by reviewing the physical, biogeochemical, and microbiological characteristics of microbial habitats that have been identified within glaciers and ice sheets. Two key glacial ecosystems emerge, one inhabiting the glacier surface (the supraglacial ecosystem) and one at the ice-bed interface (the subglacial ecosystem). The supraglacial ecosystem is characterized by a diverse consortium of microbes (usually bacteria, algae, phytoflagellates, fungi, viruses and occasional rotifers, tardigrades, and diatoms) within the snowpack, supraglacial streams, and melt pools (cryoconite holes). The subglacial system is dominated by aerobic/anaerobic bacteria and most probably viruses in basal ice/till mixtures and subglacial lakes. A third, so-called englacial ecosystem is also described, but it is demonstrated that conditions within glacier ice are sufficient to make metabolic activity and its impact upon nutrient dynamics negligible at the glacier scale.Consideration of the surface and internal heat balances of the glacier show that all glacial ecosystems are sensitive to climate change, although at different timescales. Thus, while rapid, melt-driven habitat changes lead to melt-out, resuscitation, and redistribution of microorganisms in many supraglacial ecosystems, much slower climatic and glacial mass-balance processes effect such changes in the subglacial ecosystem. Paradoxically, it is shown that these forces have brought about net refreezing and the onset of cryostasis in the subglacial ecosystems of many Arctic glaciers subject to thinning in recent decades.

Darkening of parts of the Greenland ice sheet surface during the summer months leads to reduced albedo and increased melting. Here we show that heavily pigmented, actively photosynthesising microalgae and cyanobacteria are present on the bare ice. We demonstrate the widespread abundance of green algae in the Zygnematophyceae on the ice sheet surface in Southwest Greenland. Photophysiological measurements (variable chlorophyll fluorescence) indicate that the ice algae likely use screening mechanisms to downregulate photosynthesis when exposed to high intensities of visible and ultraviolet radiation, rather than non-photochemical quenching or cell movement. Using imaging microspectrophotometry, we demonstrate that intact cells and filaments absorb light with characteristic spectral profiles across ultraviolet and visible wavelengths, whereas inorganic dust particles typical for these areas display little absorption. Our results indicate that the phototrophic community growing directly on the bare ice, through their photophysiology, most likely have an important role in changing albedo, and subsequently may impact melt rates on the ice sheet.

Cryoconite holes, which can cover 0.1-10% of the surface area of glaciers, are small, water-filled depressions (typically o1 m in diameter and usually o0.5 m deep) that form on the surface of glaciers when solar-heated inorganic and organic debris melts into the ice. Recent studies show that cryoconites are colonized by a diverse range of microorganisms, including viruses, bacteria and algae. Whether microbial communities on the surface of glaciers are actively influencing biogeochemical cycles or are just present in a dormant state has been a matter of debate for long time. Here, we report primary production and community respiration of cryoconite holes upon glaciers in Svalbard, Greenland and the European Alps. Microbial activity in cryoconite holes is high despite maximum temperatures seldom exceeding 0.1 1C. In situ primary production and respiration in cryoconites during the summer is often comparable with that found in soils in warmer and nutrient richer regions. Considering only glacier areas outside Antarctica and a conservative average cryoconite distribution on glacial surfaces, we found that on a global basis cryoconite holes have the potential to fix as much as 64 Gg of carbon per year (i.e. 98 Gg of photosynthesis minus 34 Gg of community respiration). Most lakes and rivers are generally considered as heterotrophic systems, but our results suggest that glaciers, which contain 75% of the freshwater of the planet, are largely autotrophic systems.

Abstract. This paper describes detailed budgets of water, Cl ) , dissolved Si and both inorganic and organic forms of nitrogen and phosphorus for two small glacier basins in Arctic Svalbard (Midre Love´nbreen and AustreBrøggerbreen). Rates of nutrient deposition are modest, dominated by inorganic nitrogen and episodically enhanced by extreme events. Hence deposition rates are also variable, ranging from 20 to 72 kg NO 3 -N km )2 a )1 and 10-37 kg NH 4 -N km )2 a )1 over just two consecutive years. Deposition of dissolved organic and particulate forms of nitrogen (DONand PN respectively) also appears significant and therefore requires further investigation (3-8 kg DON-N km )2 and 7-26 kg PN-N km )2 during winter -no summer data are available). Evidence for microbially mediated nutrient cycling within the glacial system is clear in the nutrient budgets, as is the release of large phosphorus, Si and organic/particulate nitrogen fluxes by subglacial erosion. The latter is entirely dependent upon the presence of subglacial drainage, promoting silicate mineral dissolution and the erosion of largely unweathered apatite. The large DON and PN fluxes are surprising and may relate to young organic nitrogen associated with microbial life within the glaciers. This is because wide spread assimilation of NH 4 + and perhaps even nitrification occurs on the glacier surface, most likely within abundant cryoconite holes. Further microbial activity also occurs at the glacier bed, where denitrification and sulphate reduction is now known to take place. Thus a two component 'glacial ecosystem' is proposed that is highly sensitive to climate change.

[1] The manner by which meltwater drains through a glacier is critical to ice dynamics, runoff characteristics, and water quality. However, much of the contemporary knowledge relating to glacier hydrology has been based upon, and conditioned by, understanding gleaned from temperate valley glaciers. Globally, a significant proportion of glaciers and ice sheets exhibit nontemperate thermal regimes. The recent, growing concern over the future response of polar glaciers and ice sheets to forecasts of a warming climate and lengthening summer melt season necessitates recognition of the hydrological processes in these nontemperate ice masses. It is therefore timely to present an accessible review of the scientific progress in glacial hydrology where nontemperate conditions are dominant. This review provides an appraisal of the glaciological literature from nontemperate glaciers, examining supraglacial, englacial, and subglacial environments in sequence and their role in hydrological processes within glacierized catchments. In particular, the variability and complexity in glacier thermal regimes are discussed, illustrating how a unified model of drainage architecture is likely to remain elusive due to structural controls on the presence of water. Cold ice near glacier surfaces may reduce meltwater flux into the glacier interior, but observations suggest that the transient thermal layer of near surface ice holds a hydrological role as a depth-limited aquifer. Englacial flowpaths may arise from the deep incision of supraglacial streams or the propagation of hydrofractures, forms which are readily able to handle varied meltwater discharge or act as locations for water storage, and result in spatially discrete delivery of water to the subglacial environment. The influence of such drainage routes on seasonal meltwater release is explored, with reference to summer season upwellings and winter icing formation. Moreover, clear analogies emerge between nontemperate valley glacier and ice sheet hydrology, the discussion of which indicates how persistent reassessment of our conceptualization of glacier drainage systems is required. There is a clear emphasis that continued, integrated endeavors focused on process glaciology at nontemperate glaciers are a scientific imperative to augmenting the existing body of research centered on ice mass hydrology.

[1] This paper presents the results of a distributed, two-dimensional surface energy balance model used to investigate the spatial and temporal variations in the surface energy balance of Midre Lovénbreen, a small valley glacier in northwest Spitsbergen, Svalbard, over the summer of 2000. We utilize high-resolution airborne lidar data to derive a digital elevation model of the glacier and surrounding topography, on which a surface energy balance is computed, driven by meteorological data obtained from a meteorological station located on the glacier and a synoptic station maintained at the nearby Ny-Å lesund research base. Given the high-resolution topographic data, we focus particularly on whether the long duration of sunshine at high latitudes compensates for the higher solar zenith angles on the season-long energy balance and whether shading by the surrounding topography plus glacier surface slope and aspect play an increased role in the patterns of solar radiation receipt (and hence melt) over the glacier surface. The model results are validated using a combination of mass balance data from the glacier, measured surface lowering at the glacier meteorological station, and by comparing a derived energy balance component from the model with a measured energy flux. Overall, the model performance is very good. Glacier topography is found to play a fundamental role in determining the surface energy balance; topographic shading, slope, and aspect and correction of the surface albedo for high solar zenith angles are found to play a crucial role in determining spatial patterns of surface energy balance and therefore melt.

[1] The basal regions of continental ice sheets are gaps in our current understanding of the Earth's biosphere and biogeochemical cycles. We draw on existing and new chemical data sets for subglacial meltwaters to provide the first comprehensive assessment of sub-ice sheet biogeochemical weathering. We show that size of the ice mass is a critical control on the balance of chemical weathering processes and that microbial activity is ubiquitous in driving dissolution. Carbonate dissolution fueled by sulfide oxidation and microbial CO 2 dominate beneath small valley glaciers. Prolonged meltwater residence times and greater isolation characteristic of ice sheets lead to the development of anoxia and enhanced silicate dissolution due to calcite saturation. We show that sub-ice sheet environments are highly geochemically reactive and should be considered in regional and global solute budgets. For example, calculated solute fluxes from Antarctica (72-130 t yr −1 ) are the same order of magnitude as those from some of the world's largest rivers and rates of chemical weathering (10-17 t km −2 yr −1 ) are high for the annual specific discharge (2.3-4.1 × 10 −3 m). Our model of chemical weathering dynamics provides important information on subglacial biodiversity and global biogeochemical cycles and may be used to design strategies for the first sampling of Antarctic Subglacial Lakes and other sub-ice sheet environments for the next decade.

Glacier surfaces support unique microbial food webs dominated by organic and inorganic debris called 'cryoconite'. Observations from Longyearbreen, Spitsbergen, show how these aggregate particles can develop an internal structure following the cementation of mineral grains (mostly quartz and dolomite) by filamentous microorganisms. Measurements of carbon and dissolved O2 show that these microorganisms, mostly cyanobacteria, promote significant rates of photosynthesis (average 17 ?gC g?1 d?1) which assist aggregate growth by increasing the biomass and producing glue-like extracellular polymeric substances. The primary production takes place not only upon the surface of the aggregates but also just beneath, due to the translucence of the quartz particles. However, since total photosynthesis is matched by respiration (average 19 ?gC g?1 d?1), primary production does not contribute directly to cryoconite accumulation upon the glacier surface. The microorganisms therefore influence the surface albedo most by cementing dark particles and organic debris together, rather than simply growing over it. Time-lapse photographs show that cryoconite is likely to reside upon the glacier for years as a result of this aggregation. These observations therefore show that a better understanding of the relationship between supraglacial debris and ablation upon glaciers requires an appreciation of the biological processes that take place during summer.Peer reviewe